Behavioural interference between ungulate species: roe are not on velvet with fallow deer

Interference is expected to occur at feeding areas between species with a similar diet, but few studies have tested this idea for wild ungulates. We analysed interactions between fallow deer, European roe deer and wild boar, in three sites, in a Mediterranean area. We expected that interference should be greater between deer than between them and wild boar. We documented the negative effects of behavioural interference by fallow on foraging behaviour of roe deer, under field conditions. Deer species built up 90% interference interactions, with fallow always dominant on roe, also through direct aggression. Although roe deer decreased feeding and increased vigilance levels in proximity (<50 m) of either fallow deer or wild boar, they were displaced significantly more often by the former than by the latter. Fallow deer were neither displaced nor alarmed by roe and rarely by wild boar. No deer species displaced wild boar. Interference was significantly greater on solitary roe deer, especially females, in spring and roe left the feeding ground most often in the smallest site (13 ha). Roe deer avoided areas where the local density of fallow deer was the highest. During our 4-year-study, roe deer density decreased whereas fallow deer numbers increased. Behavioural interference may explain how fallow deer outcompete roe deer through spatial exclusion from feeding sites and avoidance of areas with high densities of the former. Fallow deer evolved in semi-arid, relatively poor habitats of Asia Minor: interspecific defence of crucial resources could have developed as a beneficial tactic for its survival.     

Forests Too Deer: Edge Effects in Northern Wisconsin

Abstract: Browsing by white-tailed deer (Odocoileus virginianus) can profoundly affect the abundance and population structure of several woody and herbaceous plant species. Enclosure studies and population surveys reveal that past and current deer densities as low as 4 deer/km² may prevent regeneration of the once common woody species, Canada yew (Taxus canadensis), eastern hemlock (Tsuja canadensis), and white cedar Puja occidentalis), as well as several herbaceous species. Prior to European settlement, forests in northern Wisconsin contained relatively sparse deer populations (<4/km²), but extensive timber cutting in the late nineteenth century boosted deer populations. Continued habitat fragmentation resulting from scattered timber harvests and the creation of "wildlife openings" to improve deer forage maintain these high densities throughout much of the Northeast.
Because deer wander widely, the effects of high deer densities penetrate deeply into remaining stands of old and mature forest, greatly modifying their composition Thus, abundant early successional and "edge" habitat, and the high deer densities they engender, represent significant external threats to these plant communities. We hypothesize that establishing large (200–400 km²) continuous areas of maturing forest, especially in conjunction with increased hunting, could reduce local deer densities and so provide a simple and inexpensive method for retaining species sensitive to the deleterious effects of browsing.     

The effect of deer management on the abundance of Ixodes ricinus in Scotland

The management of wildlife hosts for controlling parasites and disease has a history of mixed success. Deer can be important hosts for ticks, such as Ixodes ricinus, which is the primary vector of disease-causing zoonotic pathogens in Europe. Deer are generally managed by culling and fencing for forestry protection, habitat conservation, and commercial hunting, and in this study we test whether these deer management methods can be useful for controlling ticks, with implications for tick-borne pathogens. At different spatial scales and habitats we tested the hypotheses that tick abundance is reduced by (1) culling deer and (2) deer exclusion using fencing. We compared abundance indices of hosts and questing I. ricinus nymphs using a combination of small-scale fencing experiments on moorland, a large-scale natural experiment of fenced and unfenced pairs of forests, and cross-sectional surveys of forest and moorland areas with varying deer densities. As predicted, areas with fewer deer had fewer ticks, and fenced exclosures had dramatically fewer ticks in both large-scale forest and small-scale moorland plots. Fencing and reducing deer density were also associated with higher ground vegetation. The implications of these results on other hosts, pathogen prevalence, and disease risk are discussed. This study provides evidence of how traditional management methods of a keystone species can reduce a generalist parasite, with implications for disease risk mitigation.     

一种用粪便样方法估计林麝种群密度的新方法

数量(密度)特征是动物种群的一个重要特征，是定点研究一个野生种群生态的一项非常重要的基础工作．其数量和年龄结构为进一步分析其种群的密度、移动、巢域和核域，尤其是为种群动态提供依据，也是珍稀动物保护和资源动物管理利用的先行工作和重要组成部份．关于野生动物种群数量的具体统计方法，有不少的文章报道，然而从动物的空间分布格局来研究大型脊椎动物却较为少见，这主要是受研究方法的限制．为此，作者对此进行了改进，提出了一种新的研究方法：临时空间样方法，并将其应用到冶勒自然保护区林麝的数量调查研究中去．结果表明：(1)集聚分布指数CA=12．09，说明冶勒自然保护区的林麝有较高的集聚度；(2)林麝粪堆在野外的分布属于聚集分布，并与负二项分布拟合较好；(3)在冶勒自然保护区，生活有62～98只左右的林麝，其密度为：1．0～1．6只／km^2．林麝在冶勒自然保护区呈现聚积分布的原因是由于环境和人类活动因素造成的．图1表2参24     

Emerging prion disease drives host selection in a wildlife population

Infectious diseases are increasingly recognized as an important force driving population dynamics, conservation biology, and natural selection in wildlife populations. Infectious agents have been implicated in the decline of small or endangered populations and may act to constrain population size, distribution, growth rates, or migration patterns. Further, diseases may provide selective pressures that shape the genetic diversity of populations or species. Thus, understanding disease dynamics and selective pressures from pathogens is crucial to understanding population processes, managing wildlife diseases, and conserving biological diversity. There is ample evidence that variation in the prion protein gene (PRNP) impacts host susceptibility to prion diseases. Still, little is known about how genetic differences might influence natural selection within wildlife populations. Here we link genetic variation with differential susceptibility of white-tailed deer to chronic wasting disease (CWD), with implications for fitness and disease-driven genetic selection. We developed a single nucleotide polymorphism (SNP) assay to efficiently genotype deer at the locus of interest (in the 96th codon of the PRNP gene). Then, using a Bayesian modeling approach, we found that the more susceptible genotype had over four times greater risk of CWD infection; and, once infected, deer with the resistant genotype survived 49% longer (8.25 more months). We used these epidemiological parameters in a multi-stage population matrix model to evaluate relative fitness based on genotype-specific population growth rates. The differences in disease infection and mortality rates allowed genetically resistant deer to achieve higher population growth and obtain a long-term fitness advantage, which translated into a selection coefficient of over 1% favoring the CWD-resistant genotype. This selective pressure suggests that the resistant allele could become dominant in the population within an evolutionarily short time frame. Our work provides a rare example of a quantifiable disease-driven selection process in a wildlife population, demonstrating the potential for infectious diseases to alter host populations. This will have direct bearing on the epidemiology, dynamics, and future trends in CWD transmission and spread. Understanding genotype-specific epidemiology will improve predictive models and inform management strategies for CWD-affected cervid populations.     

Modelling landscape effects on density-contact rate relationships of deer in eastern Alberta: Implications for chronic wasting disease

Managing wildlife diseases requires an understanding of disease transmission, which may be strongly affected by host population density and landscape features. Transmission models are typically fit from time-series disease prevalence data and modelled based on how the contact rate among hosts is affected by density, which is often assumed to be a linear (density-dependent transmission) or constant (frequency-dependent transmission) relationship. However, long-term time-series data is unavailable for emerging diseases, and this approach cannot account for independent effects of landscape. We developed a mechanistic model based on ecological data to empirically derive the contact rate-density relationship in white-tailed and mule deer in an enzootic region of chronic wasting disease (CWD) in Alberta, Canada and to determine whether it was affected by landscape. Using data collected from aerial surveys and GPS-telemetry, we developed empirical relationships predicting deer group size, home range size, and habitat selection to iteratively simulate deer distributions across a range of densities and landscapes. We calculated a relative measure of total per-capita contact rate, which is proportional to the number of other deer contacted per individual per unit time, for each distribution as the sum of pairwise contact rates between a target deer and all other individuals. Each pairwise contact rate was estimated from an empirical relationship developed from GPS-telemetry data predicting pairwise contact rates as a function of home range overlap and landscape structure. Total per-capita contact rates increased as a saturating function of density, supporting a transmission model intermediate between density- and frequency-dependent transmission. This pattern resulted from group sizes that reached an asymptote with increasing deer density, although this relationship was mediated by tree and shrub coverage in the landscape, such that in heavily wooded areas, the contact rate saturated at much lower densities. These results suggest that CWD management based on herd reductions, which require a density-dependent contact rate to be effective, may have variable effects on disease across a single management region. The novel mechanistic approach we employed for estimating effects of density and landscape on transmission is a powerful complement to typical data-fitting approaches for modelling disease transmission.     

Simulation and analysis of outbreaks of bark beetle infestations and their management at the stand level

Outbreaks of bark beetles in forests can result in substantial economic losses. Understanding the factors that influence the development and spread of bark beetle outbreaks is crucial for forest management and for predicting outbreak risks, especially with the expected global warming. Although much research has been done on the ecology and phenology of bark beetles, the complex interplay between beetles, host trees, beetle antagonists and forest management makes predicting beetle population development especially difficult. Using the recent infestations of the European Spruce Bark Beetle (Ips typographus L. Col. Scol.) in the Bavarian Forest National Park (Germany) as a case study, we developed a spatially explicit agent-based simulation model (SAMBIA) that takes into account individual trees and beetles. This model primarily provides a tool for analysing and understanding the spatial and temporal aspects of bark beetles outbreaks at the stand scale. Furthermore, the model should allow an estimation of the effectiveness of concurrent impacts of both antagonists and management to confine outbreak dynamics in practice. We also used the model to predict outbreak probabilities in various settings. The simulation results indicated a distinct threshold behaviour of the system in response to pressure by antagonists or management of the bark beetle population. Despite the different scenarios considered, we were able to extract from the simulations a simple rule of thumb for the successful control of an outbreak: if roughly 80% of individual beetles are killed by antagonists or foresters, outbreaks will rarely take place. Our model allows the core dynamics of this complex system to be reduced to this inherent common denominator.     

A Voronoi diagram based population model for social species of wildlife

A population model is presented that accounts for spatial structure within habitat patches. It is designed for social species of wildlife that form social group home ranges that are much smaller than patch size. The model represents social group home ranges by Voronoi regions that tessellate a patch to form a Voronoi diagram. Neighbouring social groups are linked with habitat-confined shortest paths and form a dispersal network. The model simulates population dynamics and makes use of Voronoi diagrams and dispersal networks as a spatial component. It then produces density maps as outputs. These are maps that show predicted animal densities across the patches of a landscape. A construction procedure for the particular Voronoi diagram type used by the model is described. As a test case, the model is run for the squirrel glider (Petaurus norfolcensis), a small arboreal marsupial native to Australia. A time series of density maps are produced that show squirrel glider density changing across a landscape through time.     

Modeling the population dynamics of capybara Hydrochaeris hydrochaeris: a first step towards a management plan

Capybara (Hydrochaeris hydrochaeris) is the largest living rodent, widely distributed in South America, with a considerable potential as an economic resource. There are large populations in the region of Esteros del Ibera that could be exploited in a sustainable manner and contribute positively to the development of the region. Such exploitation requires to be done responsibly and following a specially designed management plan. In the particular case of wildlife management, a sustainable exploitation of a given species requires knowledge of its dynamics, its density in the area, and its vital rates, as well as understanding how the natural environment will respond to the proposed manipulation. It is also important to determine which environmental variables lead the population dynamics in each region. A mathematical model is a very convenient tool for analyzing possible strategies.This work presents a matrix population model structured in five stages. Sensitivity and elasticity analyses of the model provide an understanding of the effects of differentiated class-dependent survival and growth probabilities on the life cycle. A harvest term has been included in the model with the purpose of evaluating the sustainability of a given exploitation scenario or comparing different harvest strategies. The different strategies depend on the exploitation pattern, the exploitation intensity, and the season in which the harvest takes place.The purpose is to formulate a simple tool that can be understood by a manager without specific training. Hence, the software interface was specially designed for a user with elementary knowledge of vital parameters of the species.     

Ungulate population dynamics and optimization models

Optimal harvesting strategies for an ungulate population are estimated using stochastic dynamic programming. Data on the Llano Basin white-tailed deer (Odocoileus virginianus) population were used to construct a 2-variable population dynamics model. The model provided the basis for estimating optimal harvesting strategies as a feedback function of the current values of the state variables (prefawning older deer and juveniles). Optimal harvest strategies were insensitive to assumptions about the probability distributions of the stochastic variable (rainfall). The response of the population components to harvesting and the returns obtained from applying optimal strategies were explored through simulation. Mean annual harvest is about 15% of the population. Simplified harvesting strategies based on age-ratios as well as a simplified version based on optimal strategies—but assuming persisting equilibrium juvenile deer density—were compared to optimal strategies through examining values of information. Simplified harvesting strategies lead to a lower harvest over a 50-year simulation period.     

Lifetime reproductive success and composition of the home range in a large herbivore

The relationship between individual performance and nonrandom use of habitat is fundamental to ecology; however, empirical tests of this relationship remain limited, especially for higher orders of selection like that of the home range. We quantified the association between lifetime reproductive success (LRS) and variables describing lifetime home ranges during the period of maternal care (spring to autumn) for 77 female roe deer (Capreolus capreolus) at Trois-Fontaines, Champagne-Ardenne, France (1976-2000). We maintained population growth rate (adjusted to account for removals of non-focal animals) near rmax, which enabled us to define the fitness-habitat relationship in the absence of density effects. Using a negative binomial model, we showed that a roe deer's incorporation into its home range of habitat components important to food, cover, and edge (meadows, thickets, and increased density of road allowances) was significantly related to LRS. Further, LRS decreased with increasing age of naturally reclaimed meadows at the time of a deer's birth, which may have reflected a cohort effect related to, but not entirely explained by, a decline in quality of meadows through time. Predictive capacity of the selected model, estimated as the median correlation (rs) between predicted and observed LRS among deer of cross-validation samples, was 0.55. The strength of this relationship suggests that processes like selection of the site of a home range during dispersal may play a more important role in determining fitness of individuals than previously thought. Individual fitness of highly sedentary income breeders with high reproductive output such as roe deer should be more dependent on home range quality during the period of maternal care compared to capital breeders with low reproductive output. Identification of the most important habitat attributes to survival and reproduction at low density (low levels of intraspecific competition) may prove useful for defining habitat value ("intrinsic habitat value").     

Influence of host migration between woodland and pasture on the population dynamics of the tick Ixodes ricinus: A modelling approach

Ticks act as vectors of pathogens that can be harmful to animals and/or humans. Epidemiological models can be useful tools to investigate the potential effects of control strategies on diseases such as tick-borne diseases. The modelling of tick population dynamics is a prerequisite to simulating tick-borne diseases and the corresponding spread of the pathogen. We have developed a dynamic model to simulate changes in tick density at different stages (egg, larva, nymph and adult) under the influence of temperature. We have focused on the tick Ixodes ricinus, which is widespread in Europe. The main processes governing the biological cycles of ticks were taken into account: egg laying, hatching, development, host (small, mainly rodents, or large, like deer and cattle, mammals) questing, feeding and mortality. This model was first applied to a homogeneous habitat, where simulations showed the ability of the model to reproduce the general patterns of tick population dynamics. We considered thereafter a multi-habitat model, where three different habitats (woodland, ecotone and meadow) were connected through host migration. Based on this second application, it appears that migration from woodland, via the ecotone, is necessary to sustain the presence of ticks in the meadow. Woodland can therefore be considered as a source of ticks for the meadow, which in turn can be regarded as a sink. The influence of woodland on surrounding tick densities increases in line with the area of this habitat before reaching a plateau. A sensitivity analysis to parameter values was carried out and demonstrated that demographic parameters (sex ratio, development, mortality during feeding and questing, host finding) played a crucial role in the determination of questing nymph densities. This type of modelling approach provides insight into the influence of spatial heterogeneity on tick population dynamics.     

Rapid declines of large mammal populations after the collapse of the Soviet Union

Anecdotal evidence suggests that socioeconomic shocks strongly affect wildlife populations, but quantitative evidence is sparse. The collapse of socialism in Russia in 1991 caused a major socioeconomic shock, including a sharp increase in poverty. We analyzed population trends of 8 large mammals in Russia from 1981 to 2010 (i.e., before and after the collapse). We hypothesized that the collapse would first cause population declines, primarily due to overexploitation, and then population increases due to adaptation of wildlife to new environments following the collapse. The long‐term Database of the Russian Federal Agency of Game Mammal Monitoring, consisting of up to 50,000 transects that are monitored annually, provided an exceptional data set for investigating these population trends. Three species showed strong declines in population growth rates in the decade following the collapse, while grey wolf (Canis lupus) increased by more than 150%. After 2000 some trends reversed. For example, roe deer (Capreolus spp.) abundance in 2010 was the highest of any period in our study. Likely reasons for the population declines in the 1990s include poaching and the erosion of wildlife protection enforcement. The rapid increase of the grey wolf populations is likely due to the cessation of governmental population control. In general, the widespread declines in wildlife populations after the collapse of the Soviet Union highlight the magnitude of the effects that socioeconomic shocks can have on wildlife populations and the possible need for special conservation efforts during such times. Declinación Rápida de las Poblaciones de Mamíferos Mayores después del Colapso de la Unión Soviética     

An agent-based model of red colobus resources and disease dynamics implicates key resource sites as hot spots of disease transmission

The effect of anthropogenic landscape change on disease in wildlife populations represents a growing conservation and public health concern. Red colobus monkeys (Procolobus rufomitratus), an endangered primate species, are particularly susceptible to habitat alteration and have been the focus of a great deal of disease and ecological research as a result. To infer how landscape changes can affect host and parasite dynamics, a spatially explicit agent-based model is created to simulate movement and foraging of this primate, based on a resource landscape estimated from extensive plot-derived tree population data from Kibale National Park, Uganda. Changes to this resource landscape are used to simulate effects of anthropogenic forest change. With each change in the landscape, disease outcomes within the simulated red colobus population are monitored using a hypothetical microparasite with a directly transmitted life cycle. The model predicts an optimal distribution of resources which facilitates the spread of an infectious agent through the simulated population. The density of resource rich sites and the overall heterogeneity of the landscape are important factors contributing to this spread. The characteristics of this optimal distribution are similar to those of logged sections of forest adjacent to our study area.     

Development and validation of an individual based Daphnia magna population model: The influence of crowding on population dynamics

An individual-based model was developed to predict the population dynamics of Daphnia magna at laboratory conditions from individual life-history traits observed in experiments with different feeding conditions. Within the model, each daphnid passes its individual life cycle including feeding on algae, aging, growing, developing and – when maturity is reached – reproducing. The modelled life cycle is driven by the amount of ingested algae and the density of the Daphnia population. At low algae densities the population dynamics is mainly driven by food supply, when the densities of algae are high, the limiting factor is “crowding” (a density-dependent mechanism due to chemical substances released by the organisms or physical contact, but independent of food competition).     

Longleaf pine (Pinus palustris) and hardwood dynamics in a fire-maintained ecosystem: A simulation approach

Longleaf pine (Pinus palustris) savannas of the southeastern U.S. represent an archetype of a fire dependent ecosystem. They are known to have very short fire return intervals (∼1-3 years) that perpetuate understory plant diversity (up to 50 species m⁻²), support pine recruitment, and suppress fire sensitive hardwoods. Understanding the relationships that regulate longleaf and southern hardwoods is especially critical. With decreased fire frequency, insufficient intensity, or lack of underground competition, a woody mid-story rapidly develops, dominated by fire sensitive trees and shrubs that in-turn suppress more fire dependent species (including pine seedlings). This may occur in forest gaps, where pine-needle abundance is diminished, reducing fire spread potential. The interactions between longleaf pine, hardwoods, forest fuels, and fire frequency are complex and difficult to understand spatially. The objective of this study was to develop a spatially explicit longleaf pine-hardwood stochastic simulation model (LLM), incorporating tree demography, plant competition, and fuel and fire characteristics. Data from two longleaf pine study sites were used to develop and evaluate the model with the goal to incorporate simple site-specific calibration parameters for model versatility. Specific model components included pine seed masting, hardwood clonal sprouting, response to fire (re-sprouting, mortality), and tree density driven competition effects. LLM spatial outputs were consistent with observed forest gap dynamics associated with pine seedling establishment and hardwood encroachment. Changes in fire frequency (i.e., fire probability = 0.35-0.05) illustrated a shift in community structure from longleaf pine dominated to a hardwood dominated community. This approach to assessing model response may be useful in characterizing longleaf ecosystem resilience, especially at intermediate fire frequencies (e.g., 0.15) where the community may be sensitive to small changes in the fire regime. Height distributions and population densities were similar to in situ findings (field and LIDAR data) for both study sites. Height distributions output by the LLM illustrated fluctuations in population structure. The LLM was especially useful in determining knowledge gaps associated with fuel and fire heterogeneity, plant-plant interactions, population structure and its temporal fluctuations, and hardwood demography. This is the first known modeling work to simulate interactions between longleaf pine and hardwoods and provides a foundation for further studies on fire and forest management, especially in relation to ecological forestry practices, restoration, and site-specific applications.     

Polychlorinated biphenyl (PCB) residues in European roe deer (Capreolus capreolus) and red deer (Cervus elaphus) from north-western Poland

The purpose of this study was to evaluate polychlorinated biphenyls (PCBs) contamination levels in roe and red deer from north-western Poland and to assess environmental pollution in this area. A quantitative analysis was conducted using a capillary gas chromatography/mass spectrometry method. The mean concentrations of ΣPCBs (sum of PCBs: 28, 52, 101, 138, 153, 180) in liver samples were 30.24±12.35 ng·g^?1 of lipid weight (l.w.) in roe deer and 60.13±14.23 ng·g^?1 l.w. in red deer, compared with 24.21±10.02 and 45.22±9.77 ng·g^?1 in the lungs of roe and red deer, respectively. PCBs 138, 153 and 180 were the dominant congeners in the liver samples of the analysed animals, whereas PCB 138 and 153 in the lungs. TEQs levels calculated for only dioxin-like PCBs were low: 0.32 and 0.29 pg WHO-PCB-TEQ·g^?1 fat in liver of red deer and roe deer, respectively. The mean PCB concentrations obtained in our study for organs of roe deer and red deer were several times lower than those reported elsewhere. These findings show that the investigated roe and red deer originated from an area with low levels of PCB contamination.     

Managing the Abundance and Diversity of Breeding Bird Populations through Manipulation of Deer Populations

Abstract: Deer densities in forests of eastern North America are thought to have significant effects on the abundance and diversity of forest birds through the role deer play in structuring forest understories. We tested the ability of deer to affect forest bird populations by monitoring the density and diversity of vegetation and birds for 9 years at eight 4-ha sites in northern Virginia, four of which were fenced to exclude deer. Both the density and diversity of understory woody plants increased following deer exclosure. The numerical response of the shrubs to deer exclosure was significantly predicted by the soil quality (ratio of organic carbon to nitrogen) at the sites. Bird populations as a whole increased following exclosure of deer, particularly for ground and intermediate canopy species. The diversity of birds did not increase significantly following exclosure of deer, however, primarily because of replacement of species as understory vegetation proceeded through successional processes. Changes in understory vegetation accounted for most of the variability seen in the abundance and diversity of bird populations. Populations of deer in protected areas are capable of causing significant shifts in the composition and abundance of bird communities. These shifts can be reversed by increasing the density and diversity of understory vegetation, which can be brought about by reducing deer density.     

Can we measure carrying capacity with foraging behavior?

Carrying capacity is one of the most important, yet least understood and rarely estimated, parameters in population management and modeling. A simple behavioral metric of carrying capacity would advance theory, conservation, and management of biological populations. Such a metric should be possible because behavior is finely attuned to variation in environment including population density. We connect optimal foraging theory with population dynamics and life history to develop a simple model that predicts this sort of adaptive density-dependent change in food consumption. We then confirm the model's unexpected and manifold predictions with field experiments. The theory predicts reproductive thresholds that alter the marginal value of energy as well as the value of time. Both effects cause a pronounced discontinuity in quitting-harvest rate that we revealed with foraging experiments. Red-backed voles maintained across a range of high densities foraged at a lower density-dependent rate than the same animals exposed to low-density treatments. The change in harvest rate is diagnostic of populations that exceed their carrying capacity. Ecologists, conservation biologists, and wildlife managers may thus be able to use simple and efficient foraging experiments to estimate carrying capacity and habitat quality.